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Interactions in Soft Bottom Benthic Communities

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Interactions in Soft Bottom Benthic Communities HELGOI~NDER MEI~Rt~SUNTERSUCHUNGI~N Helgol~inder Meeresunters. 45, 301-316 (1991} Interactions in soft bottom benthic, communities: quantitative aspects of behaviour in the surface deposit feeders Pygospio elegans (Polychaeta) and Macoma balthica (Bivalvia}* Thomas Brey Alfred- Wegener-Institut ffir Polar- und Meeresforschung; ColumbusstraBe, D-W-2850 Bremerhaven, Federal Republic of Germany ABSTRACT: The surface deposit feeding species Pygospio elegans and Macoma balthica are dominant members of many sandy bottom communities of northern boreal regions. The feeding mode of both species and the tube-building of P. elegans are assumed to affect community structure by interactions with other species. The weight of tubes of P. elegans varied between 2 and 13 g DW/ 100 cm 2 at the two stations investigated and during the year, which is equivalent to 230-1500 cm of tubes per 100 cm 2 of sediment surface. Sediment stability may be affected directly or indirectly by the amount of tubes present. M. balttn'ca shows a hnear relation between the maximum size of particles which can be inhaled and animal length. In Kiel Bay, particles > 0.5 mm are out of the range of this species. In summer, the potential feeding area (PFA,) of a P. elegans population at one station in Kiel Bay was 1.8 times the available surface area. The PFA of three different populations of M. balthica in Kiel Bay exceeded the available surface area by factors of 2.6, 2.7, and 3.2. These findings indicate strong intra- and interspecific competition for food. Additionally, the feeding of both species may strongly affect the recruitment of benthic species via pelagic larvae. Experiments are proposed to evaluate the significance of the investigated behavioural aspects for community structure. INTRODUCTION Intra- and interspecific interactions and their significance for community structure in soft bottoms have been the subject of many publications during recent years (Bell & Coull, 1980; Black & Peterson, 1988; Blaricom, i982; Bonsdorff et al., 1986; Gallagher et al., 19831 Hunt et aE, 19871 Levin, 19811 Luckenbach, 1987; Olafsson, 1989; Peterson, 1979; Reise, 1983; Whiflach & Zajak, 19851 Wilson, 1983b; Woodin, 1981; and many others). Any interaction depends on the effect of one animal on another and vice versa, i.e. interactions are based on certain aspects of the life style or behaviour of the animals. In most cases, we know the mechanisms which cause a positive or negative effect of one animal on another, e.g. protection, predation, territorialism, occupation of space, or disturbance. However, with respect to soft bottom benthic communities, only a few authors have examined the "quantity" of a certain behaviour which an interaction may "AWI Publication No. 393 Biologische Anstalt Helgoland, Hamburg 302 T. Brey be based on, e.g. the movement of a meiobenthic predator (Watzin, 1985); the sediment turnover of a population of sediment feeders (Cadee, 1976 and references therein), or the pore water transport rates of a population of tube building polychaetes (Aller, 1980). The aim of this paper is to evaluate some aspects of behaviour which are potential sources of interactions in two surface deposit feeding species, the tube building polychaete t~gospio elegans (Clapar~de) and the bivalve Macoma balthica (L.), which are both very common in shallow sandy sediments of the northern boreal regions. In both species, feeding is assumed to affect other animals which hve at the sediment surface, either via disturbance and competition for food or via predation (see e.g. Hines et al., 1989; Olafsson, 1989; Wilson, 1981). I have tried to quantify the potential feeding area at the sediment surface, i.e. the area within the range of the tentacles (P. elegans) or the inhalent siphon (M. baltln'ca) of the animals. Additionally, I investigated the particle size selection of M. balthica, which may play an important role for the successful recruitment of species with pelagic larvae (see Hines et al., 1989) and the amount of sediment which is bound in the tubes of P. elegans, which are assumed to affect pore water transport and sediment stabihty. METHODS Samples were taken at two stations, the subtidal station "Gabelsflach" (GF) in Kiel Bay (medium/fine sand, 12 m water depth) and the intertidal station "Westerhever" (WH) in the German Wadden Sea {fine sand), during 1986 to 1988 {Fig. 1). Specimens for laboratory experiments were sampled at the station GF with a 0.1 m 2 VanVeen grab or a 0.09 m 2 box corer. All other samples were taken by hand {station GF: diver} operated corers (27 cm 2, 10 cm sampling depth), fixed in a seawater solution of 0.4 % formaldehyde and 3% Kohrsolin (see Brey, 1986}, stained with Bengal rose, and sieved through 0.25 mm in the laboratory. Pygospio elegans - potential feeding area (PFA) At the station GF a 40 x 30 crn PVC tray was filled with a 4 cm layer of natural sediment, which had been sieved through 1 rnm previously in order to remove larger animals. On top of this sediment layer I put unsieved sediment from the upper 3-5 cm of the content of two grabs. Afterwards, the tray was filled with seawater. After two days, the specimens of P. elegans in the tray had re-established their tubes and were easily recognizable by the area around each tube, which was swept clean of all fine detritus particles. The size of 50 (June 86) and 60 (July 87) randomly selected PFAs was measured. Pygospio elegans- tubes The amount of tubes of/9. elegans was investigated at severa] dates at both stations. The tubes were collected from core samples, dried at 80~ and weighed. In the laboratory, specimens of P. elegans were allowed to build tubes in 15-ml glass tubes filled with azoic sediment and placed in a circulating sea water system at 12 ~ Animals and their tubes were measured and weighed. 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Location of stations Gabelsflach (GF, star) in Kiei Bay and Westerhever (WH, square) in the German Wadden Sea 304 T. Brey Macoma balthica - potential feeding area (PFA) Single specimens were measured to the lower 0.1 ram, placed in a tray filled with 6 cm natural sediment and kept at 12~C. The,animals buried themselves very rapidly and started to suck material from the surface with their inhalent siphon. The area which had been swept clean was measured after 2 and 5 h in a first set of experiments, and after 24 h in a second set. I~r balthica - particle selection Self & Jumars (1988) stated that M. balthica does not select particles of a certain size with its inhalent siphon, but there may be an upper hmit of particle size which is related to the size of the animal. The maximum size of particles in the mantle cavity of preserved specimens (4-17 mm length) from the station GF was measured under the stereo microscope and correlated with animal length. RESULTS Pygospio elegans - potential feeding area (PFA) Table 1 shows the results of these experiments. The average PFA of P. elegans was 57 mm 2 {June 86) and 91 mm 2 (July 87), respectively. Figure 2 shows the frequency distribution of PFA in July 87. The minimum distance between two tubes was below 4 mm in both experiments. Direct observations showed that P. elegans is able to put the greater part of its body out of the tube, if the range of the tentacles is not sufficient to Table 1. The average feeding area of Pygospio elegans from the station GF in two laboratory experiments. Min. dist.: Minimum distance between two tubes; S.D.: Standard deviation Date N m -2 Feeding area Min. dist. Mean radius S.D. Average area (mm) (mm) (mm 2) 19 June 86 4200 3.5 4.26 0.9 57 15 July 87 3800 3,9 5.46 2.0 91 reach a certain spot at the sediment surface. Furthermore, I could not observe any sign of aggressive reactions
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